Mineral Nutrition of Livestock, 4th Edition This page intentionally left blank Mineral Nutrition of Livestock, 4th Edition Neville F. Suttle Honorary Research Fellow Moredun Foundation Pentland Science Park Bush Loan Penicuik Midlothian EH26 0PZ UK CABI is a trading name of CAB International CABI Head Offi ce CABI North American Offi ce Nosworthy Way 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org © N. Suttle 2010. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Suttle, N. F. Mineral nutrition of livestock / Neville F. Suttle. - - 4th ed. p. cm. Earlier editions entered under Eric J. Underwood. Includes bibliographical references and index. ISBN 978-1-84593-472-9 (alk. paper) 1. Minerals in animal nutrition. I. Title. SF98.M5U5 2010 636.08’527--dc22 2009022346 ISBN-13: 978 1 84593 472 9 Commissioning editor: Sarah Hulbert Production editor: Kate Hill Typeset by SPi, Pondicherry, India. Printed and bound in the UK by the MPG Books Group. Contents Preface vii 1. The Requirement for Minerals 1 2. Natural Sources of Minerals 14 3. Assessing and Controlling Mineral Status in Livestock 39 4. Calcium 54 5. Magnesium 92 6. Phosphorus 122 7. Potassium 168 8. Sodium and Chloride 182 9. Sulfur 206 10. Cobalt 223 11. Copper 255 12. Iodine 306 13. Iron 334 14. Manganese 355 15. Selenium 377 16. Zinc 426 17. Occasionally Benefi cial Elements 459 v vi Contents 18. Potentially Toxic Elements 489 19. Design of Supplementation Trials For Assessing Mineral Deprivation 528 20. Minerals and Humans 540 Appendices 555 Index 565 Preface Reviewing progress in the mineral nutrition of livestock over the last 10 years has been compli- cated by a small fraction of the many new papers that addressed practical problems in a logical way and were subjected to rigorous peer review. The focus has been either on fundamental stud- ies of molecular pathways or on commercially attractive nutritional supplements, bypassing the ‘middle-ground’ of basic mineral nutrition. Fundamental studies have rarely produced information that changes the way mineral imbalances are diagnosed, treated or avoided. Applied studies have been dominated by commercial interests and some published conclusions have been highly mis- leading, even those appearing in prestigious journals. Commercially biased experiments in min- eral nutrition can be easily recognized: experimental designs follow practices adopted by QCs during cross examinations, avoiding questions to which the answer is unknown and possibly damning; statistical significance conventions are ignored, trends (P < 0.1) becoming ‘significant’ and if that ruse fails ‘numerical superiority’ may be claimed; positive comments about supple- ments are always mentioned in the abstract, however trivial; negative comments are avoided. One journal of human clinical nutrition now appends a ‘declaration of interest’ to their published papers, with an indication of what each author has contributed to minimize biased reporting, but commercial influences are now so pervasive that a similar declaration may be needed from refe- rees. One veterinary journal now discriminates against citations of papers not subjected to peer review, a commendable practice that can greatly shorten reference lists. Rigorous review has led me to reject some widely held views, including those that unreserv- edly accredit selenomethionine and metal chelates with superior availability. While I think that Eric Underwood would have agreed with me, it seemed unfair to link his reputation to such con- tentious issues and a text which has continued to shift from the solid foundation that he so care- fully laid in 1981. Two major changes in organization have been made: first, the complete separation of calcium from phosphorus, since they are mutual antagonists rather than bedfellows when it comes to non-ruminant nutrition; second, to provide a nutritional ‘heart’ to most chapters by tagging mineral requirements behind mineral composition and availability. To counter the growing separation of mineral nutrition in man and livestock, a final chapter has been added, which highlights areas of interdependence with implications for the health of both, and the sus- tainability of their shared environment. After a decade that has seen a vast increase in the pace and complexity of research, I have been greatly assisted by the following chapter referees for covering my ignorance, particularly at the modelling and molecular levels: Professors Andrew Sykes (Lincoln College, NZ; magnesium), Susan Fairweather-Tait (University of East Anglia, UK; iron), John Arthur (Rowett Research Institute, Aberdeen; iodine and selenium) and Jerry Spears (North Carolina University; manga- nese and zinc). Old friends Bob Orskov, Dennis Scott and Chris Livesey interrupted their retire- ment, the latter’s only just begun, to put me right on the rumenology of sulfur, physiological vii viii Preface aspects of sodium and potassium and environmental aspects of potentially toxic elements, respectively. A criticism of the last edition by Professor Ivan Caple, the then Dean of University of Melbourne’s Veterinary School, that it lacked a quick reference section for busy vets, has resulted in the addition of three summary appendices, though the lists of cautionary footnotes are neces- sarily long. Readers are again invited to notify the author of errors, omissions or irrelevancies – it may be possible to respond to some of them in a subsequent paperback version. Neville Suttle Moredun Foundation Penicuik EH26 0PZ [email protected] 1 The Requirement for Minerals Early Discoveries and, early in the next century, the iodine-rich molecule thyroxine was isolated from thyroid All animal and plant tissues contain widely vary- tissue (Harington, 1926). At the beginning of ing amounts and proportions of mineral ele- the last century, it was becoming apparent that ments, which largely remain as oxides, there is more to minerals than meets the eye. carbonates, phosphates and sulfates in the ash after ignition of organic matter. In the 17th cen- tury, a Swedish chemist, Gahn, found calcium Essentiality phosphate to be the major component of bone ash. In 1875, Sir Humphrey Davy identified the Between 1928 and 1931, novel studies at element potassium in the residues of incinerated Wisconsin with rats given specially purified wood and gave it the name ‘pot ash’. The earli- diets showed that copper, manganese and zinc est hint of nutritional significance for such were each essential for health (Underwood, apparently inert substances came from Fordyce 1977). The 1930s saw the extension of such (1791), who showed that canaries on a seed studies to livestock and, in the field, animals diet required a supplement of ‘calcareous earth’ were shown to suffer from deficiencies of cop- to remain healthy and produce eggs. Calcium per and cobalt in North America, Australia and supplements were eventually used for the pre- Europe. Further studies with rats maintained in vention of rickets, a childhood disorder of bone plastic isolators to exclude atmospheric con- development that had plagued people for centu- tamination extended the list of essential miner- ries. The discovery that iron was a characteristic als to include selenium (Smith and Schwarz, component of blood led Frodisch (1832) to link 1967), an element previously renowned for its blood iron content with ‘chlorosis’ (anaemia) in toxicity to livestock. By 1981, 22 mineral ele- people. In livestock, Boussingault (1847) showed ments were believed to be essential for animal that cattle had a dietary need for common salt, life: seven major or macronutrient minerals – and Babcock (1905) induced calcium deficiency calcium, phosphorus, potassium, sodium, chlo- in dairy cows by feeding diets low in calcium. A rine, magnesium and sulfur – and 15 trace or craving of sick cattle and sheep for the bones of micronutrient mineral elements – iron, iodine, dead animals on the South African veldt led to zinc, copper, manganese, cobalt, molybdenum, the identification of phosphorus deficiency in selenium, chromium, tin, vanadium, fluorine, cattle (Theiler, 1912). Chatin (1850–1854) silicon, nickel and arsenic (Underwood, 1981). linked environmental iodine deficiency to the Subsequently, dietary supplements of alumin- incidence of endemic goitre in man and animals ium, boron, cadmium, lithium, lead and rubidium ©N. Suttle 2010. Mineral Nutrition of Livestock, 4th Edition (N. Suttle) 1 2 Chapter 1 were shown to improve growth or health in (Suzuki et al., 2005). The new millennium has rats, goats, pigs or poultry reared in highly spe- seen an explosion of activity in this field and a cialized conditions, without inducing specific new focus: the signalling mechanisms by which abnormalities or being associated with break- intracellular needs are communicated and orches- down along metabolic pathways where they trated. Calcium and superoxide ions and seleno- have specific functions (see Chapters 17 and cysteine play pivotal roles, with the selenocysteine 18); these, and some of their predecessors